THEEFFECT

OF ENZYMES
ON DIGESTION'
MICHAEL R. BEDFORD
Finnfeeds International Ltci., P.O. Box 777, Marlborough, Wilfshire, United Kingdom SN8 IXN
Phone: + 44,(0)1672 517751
F M : +44,(0)1672 517778

Primary Audience: Nutritionists, Research Scientists, Veterinarians, Feed

Formulators

differences in fiber content, content of anti-

INTRODUCTION
Despite the supposition that some minor
ingredients such as synthetic amino acids are
fully absorbed from the digestive tract, it is
highly unlikely that any major dietary ingre-
dient is ever fully digested and absorbed by
the gastrointestinal tract. The greater the
amount of undigested material, the greater the
potential for enhancement of diet digestibility.
Differences in nutrient digestibilities
exist between dietary ingredients and even
between samples of the same ingredient [l,2,
3, 41. Reasons for such observations include
nutritional factors (ANF's), processing
damage, and content of less digestible forms
(e.g. amylose vs. amylopectin) of the same
nutrient [5,6, 71.
Treatment of the diet or of individual in-
gredients with enzymes may aid in increasing
overall diet digestibility and reducing vari-
ability of ingredientsby disrupting cell walls to
allow better access of digestive enzymes to the
encapsulated nutrients, destroying ANF's,
and supplementing birds' own digestive en-
zyme array in situations when they are over-
whelmed [8, 91. A fourth mechanism which

1 Presentedat the 1996 Poultry Science Association Informal Poultry Nutrition Symposium:
"ImprovingNutrient Utilization by Ingredient and DietaryModifcation. "
 Symposium
BEDFORD 371

has only recently received attention is pro-
bably even more important than the first three
in the latter weeks of production (ie. from
3 wk of age onwards). This mechanism pro-
vides an environment which encourages
minimal bacterial fermentation in the small
intestine while encouraging beneficial bac-
terial fermentation in the caeca [lo, 111.
Although such a beneficial effect may not be
detected by a conventional faecal digestibility
assay and perhaps not even by an ileal digest-
ibility assay, the total nutrient uptake by the
bird may nevertheless be significantly en-
hanced. Thus digestibility needs careful defi-
nition when describing this effect of exogenous
enzymes.
This paper will review the most recent
IMPROVEMENT
data relating to each of the four mechanisms
INGREDIENT DIGESTIBILITY
identified and discuss the potential for future
developments in the next 5 yr.
THROUGH DESTRUCTION
IMPROVEMENT
IN ANF9s
INGREDIENTDIGESTIBILITYThe ANF’s which will be considered here
THROUGH DISRUPTION
OF include the cereal non-starch polysaccharides
PLANTCELLWALLS (NSP’s), phytic acid, and protein ANF‘s in
oilseed meals such as trypsin inhibitors and
One of the original theories put forward
to explain the success of p-glucanases in
barley-based diets was that these enzyme
complexes aided in disrupting the cell wall
structure of the endosperm, allowing more
rapid access of the birds’ endogenous amy-
lases and proteases to the cell contents [12,13].
The result would therefore be a more rapid
and complete digestion of barley. Such a result
would explain the observed improvements in
performance and AME when such enzymes
were used.
However, subsequent work demonstrated
that improvements in fat digestibility ac-
counted for a large component of the observed
response [14], and since thevast majority of fat
was not of barley origin, it stood to reason that
cell wall encapsulation could not explain all of
the response to the addition of /?-glucanases.
Simiiar effects have been noted in rye- and
wheat-based diets [15,16].
More recent work using microscopic
techniques [17] demonstrated that in wheat-
based diets there is indeed some scope for
more rapid access to endosperm cell contents
since jejunal contents of the control-fed birds
contained some undamaged endosperm cells.
Xylanase-treated birds did not contain as
much undamaged material, indicating that
cell walls were in fact being degraded by
exogenous enzymes. Whether this is a direct
(i.e. direct degradation of the cell walls) or
indirect (e.& increased disruption of cell
walls through increased gizzard/peristaltic
action from reduced intestinal viscosity) is
impossible to ascertain from the current data.
Nevertheless, this mechanism cannot be over-
looked. Comparison of the effects of the same
xylanase on six different wheat varieties indi-
cates that there is enormous variability and
that in some cases cell wall degradation by
exogenous xylanases may be minor. In others
cell wall degradation is very significant.
IN
OF
lectins.
CEREAL NSP’s
While discussion centered on cereal cell
wall encapsulation of nutrients, White et al.
[18, 191 demonstrated that the problem
elicited by barley and countered by use of
p-glucanases was in fact due to a soluble,
viscous, B-glucan component dissolved from
the endosperm cell walls; it reduced diet di-
gestibility. The beneficial effect of xylanases
in rye- and indeed wheat-based diets was the
result of a similar viscosity-reducing mecha-
nism [20]. But in the case of wheat and rye it
is arabinoxylans, not @-glucans, which are
primarily responsible for viscosity. Enzymatic
reduction of intestinal viscosity is in all cases
thought to improve nutrient digestion by re-
ducing the constraints on diffusion of all com-
ponents involved in the digestive process
(Table 1). Even relatively small reductions in
viscosity can lead to significant improvements
in nutrient digestibility [21]. Constraints on
physical convection (ie. mixing) also decline
when solution viscosity falls [22, 231. Since fat
digestion demands vigorous peristalsis to en-
sure optimum emulsification [23], it is not
 JAPR
372 EFFECT OF ENZYMES

TABLE 1. Effect of xylanase-based enzyme on ileal digestibility of nutrientsAin wheat-based diets'

surprising that the effects of viscosity reduc-
tion on fat digestion are significant. Table 2
showsthat viscosity reduction is known to have
a much greater benefit on digestion of a satu-
rated fat source such as beef tallow compared
to an unsaturated source such as soy oil [15].
The fact that digestion of tallow is more de-
pendent upon good emulsificationthan that of
the more soluble, liquid soy oil further sug-
gests that the negative effects of viscosity on
convection are considerable.
The scale of response that can be ex-
pected from enzyme use is very much de-
pendent upon the initial grain quality.
Low viscosity grains such as maize do not re-
spond markedly and consistently to viscosity-
reducing enzymes. Rye is at the other extreme:
as theviscosity increases,the bird adapts to the
perceived deficiency in digestive capacity by
increased pancreatic enzyme output and villus
surface aredgut weight [25, 261. Increased
energy utilization in the digestive process and
endogenous losses account for the reduction
in bird performance. With excessively high
viscosity, the bird has no further adaptive
capabilities, so nutrients pass through the
small intestine undigested [26].
Table 3 indicates the potential for re-
sponse and demonstrates why response to
enzyme utilization is so variable even for
one cereal grain, a result that is quite evident
(Figure 1) from data from Classen et ai. [l].
Many factors describing the quality of wheat
were determined, but only a few were found to

TABLE 2. Fat diaestibilitv and performance of birds fed tallow or soybean oil dietsA
SOY OIL SOY OIL + TALLOW TALLOW +
ENZYME ENZYME
Weight (21 days) ala 761 128' 66Sa
Feed conversion ratio (G21 days) 1.3aab 1.266b 2.449' 1.474a
Jejunal Viscosity (mPa.s) 438 32 311 139
Crude fat digestibility,% 82.3' 87.3' %.Ob 51.0'

I *Diets contained 6O%lye, 10% added fat, and a mixture of corn starch and cellulose to maintain each diet isocaloric. I
 Symposium
BEDFORD 373

CEREAL MINIMUM MAXIMUM MEAN

Maize I 15 45 2.4
IWheat I 3 I 45 I 12 I
I Triticale I 5 I 40 I 16 I
Barley 6 225 25

be relevant in describing its feeding quality
and subsequent response to enzyme addition.
It is interesting to note that the use of enzyme
addition reduces considerably the variability
between wheat samples (AME control =
31032148; + Enzyme = 3379k82) since poor
quality samples respond more than high qual-
ity wheats. Evidently, nutrient digestion was
more compromised in poor quality samples
than in high quality samples.
PHYTATE
There is an enormous amount of literature
which indicates that the use of exogenous
phytases can significantly enhance phos-
phorous digestion through destruction of
plant-derived phytates. Phytic acid is generally
regarded as being resistant to hydrolysisin the
avian gut because of the lack of any endo-
genous phytases, although this fact is not un-
questioned. Regardless of presence or ab-
sence of endogenous phytases, microbial
phytases have been shown to significantly
enhance not only phosphorous [ B ] but also
mineral [29] and protein digestion in young
chicks [B, 30,311.
The ability of phytate to create complex
proteins and digestive proteases is thought
to be responsible for its negative effect on pro-
tein digestion [32]. Hence, its destruction
will lead to a greater rate of proteolysis in
the intestinal lumen and therefore better
protein digestion. While phytate is present
in all cereal and oilseed sources to varying
degrees, the response to added phytase does
not simply rely on the concentration of its
substrate, since availabilityof phytate seems to
be markedly lower in oilseeds than in cereals,

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FIGURE 1. Influence of wheat variety and enzyme (+, with; -, without) supplementation on AME in broilers
374
for example [33]. This conclusion means that
Care must be taken in making assumptions
about the phytase susceptiblesubstrate in each
ingredient.
OILSEED MEAL, A ” s
The ANF’s of greatest interest are the
trypsin inhibitors and the lectins prevalent
in soybean meals. Diets containing any appre-
ciable quantities of these antinutrients have
been shown to be detrimental to both pig and
poultry performance and nutrient (partic-
ularly protein) digestibility [34, 35, 361.
Although high temperature processing is
well known to be effective in destroying both
of these ANFs [6, 71, consistency between
batches of soybean meal from within and es-
pecially between different processors is not
always absolute. Table 4 indicates the vari-
abdity in several quality parameters in soybean
meals identified from most parts of the world.
Evidently, even within theUnitedStates,lectin
and trypsin concentrations are not constant.
Given that 0.21 g/kg of trypsin inhibitor will
depress ileal protein digestibility of the piglet
by 15% [%I, it is evident that all of the United
States samples (if fed at 30% of the diet) would
meet or even exceed this limit.
Thus, the potential for improved animal
performance by enzymatic degradation of re-
sidual lectins and trypsin inhibitors in soybean
meal is apparent and has even been demon-
strated in rawsoya [3]. Pre-incubation of a raw
soya sample with a protease and subsequent
feeding to broilers resulted in 60% improve-
TABLE 4. Range in quality parameters of 19 soybean meal samples collected from variuos regions of the worldA
ORIGIN PROTEIN UREASE
mg N/g/min
Argentina 43.9-46.9 0.02-0.02
Brazil 46.649.2 0.02-0.15
China 43.246.1 0.02-2.59
Europe 43.449.3 0.02-0.17
India 48.249.9 0.024.22
USA 1 48.2-49.4 I 0.02-0.02
Global 1 43.2-49.9 I 0.02-259
*Data from Hessing a pl. [3].
B% of reference untoasted soybean
C~~~~~
DRaffinose + Stachyose + Verbascose
EFFECT OF ENZYMES
ment in growth rate and 23% improvement in
feed conversion ratio, compared to the raw
soya control. Consequently, the raw soya did
not significantly differ from the soybean meal
control. Moreover, modification of the soy-
bean meal process by use of lower tempera-
tures in conjunction with proteolytic enzymes
targeted at lectins and trypsin inhibitors could
potentially eliminate these factors more effec-
tively than thermal processing alone. This pro-
cess could also produce a better quality meal
since the likelihood of maillard complexing of
lysine would be reduced [37l.
IMPROVEMENT
IN
INGREDIENT
DIGESTIBILITY
THROUGH SUPPLEMENTATION
OF HOST
ENZYMES
Recent investigations into the develop-
ment of the neonatal digestive system suggest
that the digestive tract may not have sufficient
enzymatic and absorptive capacity to deal
with all types of diet [38, 39, 401. As the bird
matures, its pancreatic enzyme production
and villus surface area increase to cope with
the digestive requirements of the bird [40,41].
Almost as important, the signalling to the
pancreas becomes more mature and accurate
so that the release of enzymes from acinar
cells becomes more closely matched to the
enzymatic requirements in the intestinal
lumen, with the result that each unit of enzyme
activity is used more effectively[42]. Very little
I
 Symposi urn
BEDFORD
literature, however, is available to support
this concept. Nevertheless, work carried out
by Bedford and Classen [43]suggests that in
the first 14 days of life, performance of wheat-
soybean meal fed birds can be significantly
enhanced through augmentation of gastric
pepsin.
Such data clearly indicate that there are
circumstances when the endogenous pro-
teolytic, amylolytic, and lipolytic capacity of
the bird is exceeded, resulting in enhanced
performance though supplementation of the
host enzyme system.
IMPROVEMENT
IN
INGREDIENT
DIGESTIBILITY
THROUGH MANIPULATION OF
MICROFLORAL
POPULATIONS
Classical measurements of nutrient di-
gestibility usually focus on the faecal level, and
less frequently on the ileal level. The assump-
tions made in both circumstances are that the
difference between feed and faecal/ileal
content of any given nutrient is the result of
digestio4absorption by the bird. This assump-
tion has always been known to be subject to
microfloral interference, but this interference
was assumed to be minimal and constant,
particularly in the case of ileal measurements.
Recently, however, an increasing body of evi-
dence has questioned whether the relationship
between AME and the productive capacity or
resultant feed conversionratio of the same diet
are in fact always constant [11,44,45]. It seems
quite certain that the diet can have a signifi-
cant influence on the fermentation capacity of
microbes present in both the ileum and cae-
cum [15]and that this effect can significantly
alter the proportion of digested energy ulti-
mately utilized by the microbial population
rather than by the host. Moreover, the fact that
many of the products of microbial fermenta-
tion are still present in the faeces but volatile,
e.g. volatile fatty acids (WAS), means that
typical drying procedures used in AMERME
procedures will drive these compounds off
and result in erroneously high values.
Use of xylanolytic enzymes has been
shown to significantly alter the fermentation
profiles not only in the ileum but also the
caecum. Overall fermentation is inhibited in
the small intestine but encouraged in the
375
caeca. Reduction of ileal fermentation could
be beneficial since it is evident that the
material being fermented in this region is
predominantly undigested starch and pro-
tein, which would ordinarily be available to the
bird [ll].Stimulation of caecal fermentation
seems likely to be the result of small molecular
weight oligosaccharides coming from xylanase
action on its substrate. These oligomers can
presumably enter the caeca and provide a fer-
mentation source for caecal microbes.
Apajalahti and Morgan [&I indicate that
it is a stimulation of propionic acid production
which is responsible for the bulk of the eleva-
tion of caecal VFA's. Since this VFA is the
digestion product of fiber ordinarily not di-
gested, the net result is an energetic benefit to
the bird if it is absorbed and utilized. With
propionic acid being gluconeogenic, it is also
presumably the most valuable VFA to produce
from fiber sources. Previous work has shown
many times the effect of enzymes on fiber di-
gestion in the chick [47], but this benefit has
usually been assumed to be of little nutritional
value and will not identify the VFA pathway.
Production of lactate in the ileum on uti-
lization of Trichoderma xylanases in wheat-
based diets and increased propionate in the
caeca can also promote better gut health, since
lactic acid bacteria provide part of the basis
of competitive exclusion [a], and propionate
is identified as being toxic to salmonella
and other putrefactive bacteria. Such a shift
in microbial populations may change the
immune challenge to the bird and enhance
performance.
Thus it seems likely that the use of en-
zymes may actually increase the digestibility
and utilization of fiber through preparing it for
microbial fermentation into VFA's in the
caeca, where it may be available for chick ab-
sorption/utilization. Moreover, the shift of
bacterial metabolism from the ileum to the
caecum suggests that the current Trichoderma
xylanases increase nutrient utilization by the
host by reducing bacterial competition for the
available substrates in the small intestine.
Thus, 85% starch digestibility at the terminal
ileum in the presence of a xylanase may actu-
ally be made up of 70% due to the host and
15% due to the microflora. The situation may
be 60% and 25% in the absence of the enzyme.
If this microfloral effect were real and consis-
tent, then the greatest effect of xylanase on

376
total starch digestibilitywould be expected in
the anterior parts of the intestinal tract where
the microbial populations are minimal. This
is in fact what has been demonstrated in
chicks [U] and pigs [49], with the benefit due
to enzyme diminishing as the samples are
taken from more and more distal portions on
the gut.
In summary, nutrient digestibility needs
very careful definition when discussing re-
sponse to enzyme addition. Because of the
interaction between enzymes and the
microflora, it is evident that faecal and per-
haps even iled digestibilities may not accu-
rately reflect the proportion of nutrients
intended for use by the bird.
Much of what has been discussed relates
to intestinal viscosity. It is important to note
that intestinal viscosity is dependent upon
many interacting factors. Table 5 attempts to
TABLE 5. Factors effectinaintestinal viscosity
~~~
FACXOR
Grainvariety
Iaimate of uroduction of erain
~~~~~ ~
Processing of diet
Inclusion level of erain
Breed of bird
Age of bird
RESPONSE DIRECTION O F RESPONSE WlTH
INCREASED VISCOSITY
Proteidstarch digestibility
Fat digestibility
Microbial competition for
ileal nutrients
Microbial provision of energy
due to caecal W A ' s
JAPR
EFFECT OF ENZYMES
identify the factors suspected of being
most relevant in producing a given intestinal
viscosity.
Agken htesthdviscositymay or maynot
be detrimental to performance, depending
upon the prevalence of risk factors which re-
spond to increased viscosity. Thus, an ileal
viscosity value in mPa.s is relatively meaning-
less unless seen in context with the risk factors
prevalent at the time of viscosity measure-
ment. Table 6 attempts to identify those risk
factors which will augment the negative effects
of high viscosity.
Undoubtedly, as more work is completed
and a greater understanding of the impli-
cations of enzymes on microfloral populations
is achieved, the relationship between enzymes
and antibiotics/growthpromoters will become
more apparent. Then researchers can com-
bine these substances more effectively.
EFFECT ON VISCOSITY
I +I- I
I +I- I
Higher temperature and shear increase viscosity
Hieher inclusion levels increase viscositv I201
Breeds differ in intestinal Viscosity on identical diets [SO]
Viscosity fallswith age [Sl]
CONSIDERATIONS
Negative More apparent in younger birds when
dieestive cauacitv is limited
Negative Risk greatest with older birds since fat
level increases and more saturated fats
used [ 11,151
Increased Risk greatest in older birds with more
mature flora and in "dirty" environments
Depressed Older birds with more mature caeca
more sensitive. Use of enzymes provides
oligosaccharides for fermentation
 Symposium
BEDFORD 377

CONCLUSIONS
AND APPLICATIONS
1. The chicken is often compromised in its digestive capacity such that addition of exogenous
enzymes can improve productive performance.
2. Exogenous enzymes can improve digestion by augmenting the chick's own capacities for
protein, starch, and fat digestion, by removing ANF's which interfere with the normal
processes of digestion, or by digestion of fiber components that would otherwise pass
undigested throughout the gastrointestinal tract.
3. Interaction of the microflora in both the small intestine and caecum with the digesta makes
determination of the accurate feeding value of a fiber-degrading enzyme particularly
difficult to assess by classical digestibility techniques.
4. Exogenous enzymes may in the future be seen to play a significant role not only in animal
nutrition but most certainly in digestive tract health.

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